Confirmation of the southern African distribution of the marine sponge Hymeniacidon perlevis (Montagu, 1814) in the context of its global dispersal.

Background: Intertidal rocky shore surveys along the South African coastline (∼3,000 km) have demonstrated the presence and abundance of the encrusting orange sponge Hymeniacidon perlevis (Montagu, 1814), a well-known globally distributed species. After analysing the southern African populations, we gained a better understanding of the genetic structure of this now-accepted global species. Apart from confirming the presence of a single population of H. perlevis, we also determined its distribution in the southern African intertidal rocky shore ecosystem, compared its genetic diversity to congeners, predict its global distribution via environmental niche modelling, and discussed possible underlying mechanisms controlling the species' global distribution. Methods: We surveyed the South African coastline and sampled sponges at 53 rocky shore sites spanning over 3,000 km, from Grosse Bucht south of Lüderitz (Namibia) to Kosi Bay on the east coast of South Africa. DNA sequences of the nuclear rDNA internal transcribed spacer (ITS1) and the COI mitochondrial gene were obtained from 61 samples and compared them to a world-wide sample of other H. perlevis sequences. Using environmental predictor variables from the global dataset BIO-ORACLE, we predicted the probability of global occurrence of the species using an ensemble of eight distribution models. Results: South African specimens were found to be 99-100% identical to other populations of H. perlevis (=H. sinapium) from other world-wide regions. The presence of a single population of H. perlevis in southern Africa is supported by genetic data, extending its distribution to a relatively wide geographical range spanning more than 4,000 km along the temperate southern African coast. The predicted global occurrence by ensemble model matched well with the observed distribution. Surface temperature mean and range were the most important predictor variables. Conclusion: While H. perlevis appears to have been introduced in many parts of the world, its origins in Europe and southern Africa are unclear.

Trade-offs between bycatch and target catches in static versus dynamic fishery closures.

While there have been recent improvements in reducing bycatch in many fisheries, bycatch remains a threat for numerous species around the globe. Static spatial and temporal closures are used in many places as a tool to reduce bycatch. However, their effectiveness in achieving this goal is uncertain, particularly for highly mobile species. We evaluated evidence for the effects of temporal, static, and dynamic area closures on the bycatch and target catch of 15 fisheries around the world. Assuming perfect knowledge of where the catch and bycatch occurs and a closure of 30% of the fishing area, we found that dynamic area closures could reduce bycatch by an average of 57% without sacrificing catch of target species, compared to 16% reductions in bycatch achievable by static closures. The degree of bycatch reduction achievable for a certain quantity of target catch was related to the correlation in space and time between target and bycatch species. If the correlation was high, it was harder to find an area to reduce bycatch without sacrificing catch of target species. If the goal of spatial closures is to reduce bycatch, our results suggest that dynamic management provides substantially better outcomes than classic static marine area closures. The use of dynamic ocean management might be difficult to implement and enforce in many regions. Nevertheless, dynamic approaches will be increasingly valuable as climate change drives species and fisheries into new habitats or extended ranges, altering species-fishery interactions and underscoring the need for more responsive and flexible regulatory mechanisms.

ROV assessment of mesophotic fish and associated habitats across the continental shelf of the Amathole region.

Understanding how fish associate with habitats across marine landscapes is crucial to developing effective marine spatial planning (MSP) in an expanding and diversifying ocean economy. Globally, anthropogenic pressures impact the barely understood temperate mesophotic ecosystems and South Africa's remote Amathole shelf is no exception. The Kei and East London region encompass three coastal marine protected areas (MPAs), two of which were recently extended to the shelf-edge. The strong Agulhas current (exceeding 3 m/s), which runs along the narrow shelf exacerbates sampling challenges. For the first time, a remotely operated vehicle (ROV) surveyed fish and their associated habitats across the shelf. Results indicated fish assemblages differed between the two principle sampling areas, and across the shelf. The number of distinct fish assemblages was higher inshore and on the shelf-edge, relative to the mid-shelf. However, the mid-shelf had the highest species richness. Unique visuals of rare Rhinobatos ocellatus (Speckled guitarfish) and shoaling Polyprion americanus (wreckfish) were collected. Visual evidence of rhodolith beds, deep-water lace corals and critically endangered endemic seabreams were ecologically important observations. The ROV enabled in situ sampling without damaging sensitive habitats or extracting fish. This study provided information that supported the Amathole MPA expansions, which extended protection from the coast to beyond the shelf-edge and will guide their management. The data gathered provides baseline information for future benthopelagic fish and habitat monitoring in these new MPAs.

Shallow seamounts represent speciation islands for circumglobal yellowtail Seriola lalandi.

Phenotypic plasticity in life-history traits in response to heterogeneous environments has been observed in a number of fishes. Conversely, genetic structure has recently been detected in even the most wide ranging pelagic teleost fish and shark species with massive dispersal potential, putting into question previous expectations of panmixia. Shallow oceanic seamounts are known aggregation sites for pelagic species, but their role in genetic structuring of widely distributed species remains poorly understood. The yellowtail kingfish (Seriola lalandi), a commercially valuable, circumglobal, epipelagic fish species occurs in two genetically distinct Southern Hemisphere populations (South Pacific and southern Africa) with low levels of gene-flow between the regions. Two shallow oceanic seamounts exist in the ocean basins around southern Africa; Vema and Walters Shoal in the Atlantic and Indian oceans, respectively. We analysed rare samples from these remote locations and from the South African continental shelf to assess genetic structure and population connectivity in S. lalandi and investigated life-history traits by comparing diet, age, growth and maturation among the three sites. The results suggest that yellowtail from South Africa and the two seamounts are genetically and phenotypically distinct. Rather than mere feeding oases, we postulate that these seamounts represent islands of breeding populations with site-specific adaptations.

Connectivity and population structure of albacore tuna across southeast Atlantic and southwest Indian Oceans inferred from multidisciplinary methodology.

Albacore tuna (Thunnus alalunga) is an important target of tuna fisheries in the Atlantic and Indian Oceans. The commercial catch of albacore is the highest globally among all temperate tuna species, contributing around 6% in weight to global tuna catches over the last decade. The accurate assessment and management of this heavily exploited resource requires a robust understanding of the species' biology and of the pattern of connectivity among oceanic regions, yet Indian Ocean albacore population dynamics remain poorly understood and its level of connectivity with the Atlantic Ocean population is uncertain. We analysed morphometrics and genetics of albacore (n = 1,874) in the southwest Indian (SWIO) and southeast Atlantic (SEAO) Oceans to investigate the connectivity and population structure. Furthermore, we examined the species' dispersal potential by modelling particle drift through major oceanographic features. Males appear larger than females, except in South African waters, yet the length-weight relationship only showed significant male-female difference in one region (east of Madagascar and Reunion waters). The present study produced a genetic differentiation between the southeast Atlantic and southwest Indian Oceans, supporting their demographic independence. The particle drift models suggested dispersal potential of early life stages from SWIO to SEAO and adult or sub-adult migration from SEAO to SWIO.

New Latrunculiidae (Demospongiae, Poecilosclerida) from the Agulhas ecoregion of temperate southern Africa.

Sixteen species of Latrunculiidae Topsent, 1922, belonging to the genera Latrunculia du Bocage, 1869, Strongylodesma Lévi, 1969, Cyclacanthia Samaai Kelly, 2004, Samaai Kelly, 2002, are currently known from the temperate waters of South Africa. Extensive new sponge collections from the Amathole region of South Africa revealed the existence of three new species of Tsitsikamma, T. amatholensis sp. nov., T. madiba sp. nov., and T. beukesi sp. nov., and a new species of the endemic South African genus Cyclacanthia, C. rethahofmeyri sp. nov. With the recent addition of two new species of Tsitsikamma from Algoa Bay and Tsitsikamma National Park (T. michaeli Parker-Nance, 2019; T. nguni Parker-Nance, 2019) the total number of known South African Latrunculiidae is now 20 species in four genera. Here we propose two new subgenera of Tsitsikamma, Tsitsikamma Samaai Kelly, 2002 and Clavicaulis subgen. nov., based on the morphological groups "favus" and "pedunculata" hypothesized by Parker-Nance et al. (2019). Species in the nominotypical subgenus Tsitsikamma, containing the type species, are thick encrusting to hemispherical with a rigid honeycombed choanosome, while species in the new subgenus Clavicaulis subgen. nov. have a purse or sac-like morphology with little choanosomal structure. Despite the obvious species-level differences in morphology, multivariate analysis based on spicule measurements (anisostyle length, discorhabd length, shaft and whorl length) was not able to distinguish between the proposed Tsitsikamma species, but separated known species T. favus Samaai Kelly, 2002, T. pedunculata Samaai Kelly, 2003, and T. scurra Samaai Kelly, 2003, from each other. Similarly, DNA barcoding of the mitochondrial COI  and the nuclear ITS of Tsitsikamma specimens failed to clearly differentiate between species, but was able to differentiate sister taxon relationships within the Latrunculiidae.

Acid-base adjustments and first evidence of denticle corrosion caused by ocean acidification conditions in a demersal shark species.

Global ocean acidification is expected to chronically lower the pH to 7.3 (>2200 µatm seawater pCO2) by the year 2300. Acute hypercapnia already occurs along the South African west and south coasts due to upwelling- and low-oxygen events, with increasing frequency. In the present project we investigated the impact of hypercapnia on the endemic demersal shark species Haploblepharus edwardsii. Specifically, we experimentally analysed acid-base regulation during acute and chronic hypercapnia, the effects of chronic hypercapnia on growth rates and on denticle structure- and composition. While H. edwardsii are physiologically well adapted to acute and chronic hypercapnia, we observed, for the first time, denticle corrosion as a result of chronic exposure. We conclude that denticle corrosion could increase denticle turnover and compromise hydrodynamics and skin protection.

Strong trans-Pacific break and local conservation units in the Galapagos shark (Carcharhinus galapagensis) revealed by genome-wide cytonuclear markers.

The application of genome-wide cytonuclear molecular data to identify management and adaptive units at various spatio-temporal levels is particularly important for overharvested large predatory organisms, often characterized by smaller, localized populations. Despite being "near threatened", current understanding of habitat use and population structure of Carcharhinus galapagensis is limited to specific areas within its distribution. We evaluated population structure and connectivity across the Pacific Ocean using genome-wide single-nucleotide polymorphisms (~7200 SNPs) and mitochondrial control region sequences (945 bp) for 229 individuals. Neutral SNPs defined at least two genetically discrete geographic groups: an East Tropical Pacific (Mexico, east and west Galapagos Islands), and another central-west Pacific (Lord Howe Island, Middleton Reef, Norfolk Island, Elizabeth Reef, Kermadec, Hawaii and Southern Africa). More fine-grade population structure was suggested using outlier SNPs: west Pacific, Hawaii, Mexico, and Galapagos. Consistently, mtDNA pairwise ΦST defined three regional stocks: east, central and west Pacific. Compared to neutral SNPs (FST = 0.023-0.035), mtDNA exhibited more divergence (ΦST = 0.258-0.539) and high overall genetic diversity (h = 0.794 ± 0.014; π = 0.004 ± 0.000), consistent with the longstanding eastern Pacific barrier between the east and central-west Pacific. Hawaiian and Southern African populations group within the west Pacific cluster. Effective population sizes were moderate/high for east/west populations (738 and 3421, respectively). Insights into the biology, connectivity, genetic diversity, and population demographics informs for improved conservation of this species, by delineating three to four conservation units across their Pacific distribution. Implementing such conservation management may be challenging, but is necessary to achieve long-term population resilience at basin and regional scales.

The winter pack-ice zone provides a sheltered but food-poor habitat for larval Antarctic krill.

A dominant Antarctic ecological paradigm suggests that winter sea ice is generally the main feeding ground for krill larvae. Observations from our winter cruise to the southwest Atlantic sector of the Southern Ocean contradict this view and present the first evidence that the pack-ice zone is a food-poor habitat for larval development. In contrast, the more open marginal ice zone provides a more favourable food environment for high larval krill growth rates. We found that complex under-ice habitats are, however, vital for larval krill when water column productivity is limited by light, by providing structures that offer protection from predators and to collect organic material released from the ice. The larvae feed on this sparse ice-associated food during the day. After sunset, they migrate into the water below the ice (upper 20 m) and drift away from the ice areas where they have previously fed. Model analyses indicate that this behaviour increases both food uptake in a patchy food environment and the likelihood of overwinter transport to areas where feeding conditions are more favourable in spring.

Metal concentrations and toxicity in South African snoek (Thyrsites atun) and yellowtail (Seriola lalandi).

The concentrations of 16 metals were assessed in snoek(Thyrsites atun; n=20) and yellowtail (Seriola lalandi; n=37) sampled from the West and South-East coasts of South Africa. Variability was observed at both small (Al, Cr, Hg, Pb, Mn and Cu) and large (As and Cu) spatial scales while inter-specific examination revealed diverse metal concentrations in snoek (Higher levels: Cr, Mn, Co, Hg and Pb) and yellowtail (higher levels: Fe and Cu). Zn, As and Hg were positively correlated with yellowtail size with no such correlations in snoek. Mean concentrations of As (0.61mg·kg-1; 0.98mg·kg-1), Cd (0.008mg·kg-1; 0.004mg·kg-1), Hg (0.27mg·kg-1; 0.16mg·kg-1) and Pb (0.009mg·kg-1; 0.005mg·kg-1) for snoek and yellowtail respectively were within regulatory guidelines. However, 10% (n=2) of snoek exceeded Hg maximum allowable limit, suggesting caution and further investigation. Overall, two meals (150g) per week of snoek or larger yellowtail (12-15kg) can be safe for human (adult) consumption.

Heavy metals in marine fish meat and consumer health: a review.

The numerous health benefits provided by fish consumption may be compromised by the presence of toxic metals and metalloids such as lead, cadmium, arsenic and mercury, which can have harmful effects on the human body if consumed in toxic quantities. The monitoring of metal concentrations in fish meat is therefore important to ensure compliance with food safety regulations and consequent consumer protection. The toxicity of these metals may be dependent on their chemical forms, which requires metal speciation processes for direct measurement of toxic metal species or the identification of prediction models in order to determine toxic metal forms from measured total metal concentrations. This review addresses various shortcomings in current knowledge and research on the accumulation of metal contaminants in commercially consumed marine fish globally and particularly in South Africa, affecting both the fishing industry as well as fish consumers.

Mercury accumulation in Yellowfin tuna (Thunnus albacares) with regards to muscle type, muscle position and fish size.

The concentrations and relationships between individual mercury species and total mercury were investigated in different muscle parts and sizes of Yellowfin tuna (Thunnus albacares). Fourteen Yellowfin tuna caught in the South Atlantic off the coast of South Africa had an average total Hg (tHg) concentration of 0.77 mg/kg wet weight. No differences were detected (p > 0.05) in tHg, MethylHg (MeHg) or inorganic Hg (iHg) accumulation among the four white muscle portions across the carcass, but both tHg and iHg were found in higher concentrations (p < 0.001) in dark muscle than white muscle. Positive linear correlations with fish weight were found for both tHg (r = 0.79, p < 0.001) and MeHg (r = 0.75, p < 0.001) concentrations. A prediction model was formulated to calculate toxic MeHg concentrations from measured tHg concentrations and fish weight (cMeHg = 0.073 + 1.365 · tHg-0.008 · w). As sampling sites and subsampling methods could affect toxicity measurements, we provide recommendations for sampling guidelines.

Heavy metal accumulation and toxicity in smoothhound (Mustelus mustelus) shark from Langebaan Lagoon, South Africa.

Together with several health benefits, fish meat could introduce toxins to consumers in the form of heavy metal contaminants. High levels of mercury (Hg), especially, are frequently detected in certain predatory fish species. Mustelus mustelus fillets were analysed for 16 metals and three individual Hg species (inorganic Hg, ehtylmercury, methylmercury) with inductively coupled plasma mass spectrometry (ICP-MS) and HPLC-ICP-MS respectively. Eleven of the 30 sharks had total Hg levels above the maximum allowable limit with toxic methylmercury found as the dominant mercury species with a strong correlation (r = 0.97; p < 0.001) to total mercury concentrations. Limited correlations between metals and shark size parameters were observed; therefore metal accumulation in M. mustelus is mostly independent of size/age. Average values for arsenic (28.31 ± 18.79 mg/kg) exceed regulatory maximum limits and Hg (0.96 ± 0.69 mg/kg) is close to the maximum limit with all other metals well below maximum limits.

Marine protected area improves yield without disadvantaging fishers.

Potential fishery benefits of Marine Protected Areas (MPAs) are widely acknowledged, yet seldom demonstrated, as fishery data series that straddle MPA establishment are seldom available. Here we postulate, based on a 15-year time series of nation-wide, spatially referenced catch and effort data, that the establishment of the Goukamma MPA (18 km alongshore; 40 km²) benefited the adjacent fishery for roman (Chrysoblephus laticeps), a South African endemic seabream. Roman-directed catch-per-unit-effort (CPUE) in the vicinity of the new MPA immediately increased, contradicting trends across this species' distribution. The increase continued after 5 years, the time lag expected for larval export, effectively doubling the pre-MPA CPUE after 10 years. We find no indication that establishing the MPA caused a systematic drop in total catch or increased travel distances for the fleet. Our results provide rare empirical evidence of rapidly increasing catch rates after MPA implementation without measurable disadvantages for fishers.

The effects of gender, size and life-cycle stage on the chemical composition of smoothhound shark (Mustelus mustelus) meat.

BACKGROUND: Knowledge on the chemical composition of foods, and on the variation in such components as influenced by endogenous factors, is required for nutritional labelling, dietetic planning and food processing. This study examined the effects of endogenous factors (gender, size and life-cycle stage) on the chemical components (proximate, amino acid, fatty acid, mineral and mercury contents) of smoothhound shark (Mustelus mustelus) meat. RESULTS: Levels of moisture, protein, lipids, ash, amino acids and most minerals were not affected (P > 0.05) by the gender, size or life-cycle stages of the sharks. The fatty acid content was higher in females than in males, as well as in non-pregnant females compared to pregnant females. Certain fatty acids decreased in quantity in medium-sized sharks (prior to maturity). Pregnant females had higher aluminium (Al) and copper (Cu) levels than non-pregnant females and mercury (Hg) levels were highest in large male sharks. CONCLUSION: This study has, for the first time, established that endogenous factors have minimal influence on the chemical composition of M. mustelus meat, demonstrating the feasibility of describing an average composition for this species. Additionally, new insights have been obtained on potential safety concerns (mercury) associated with consumption of the meat that require further investigation.

Crossing invisible boundaries: the effectiveness of the Langebaan Lagoon marine protected area as a harvest refuge for a migratory fish species in South Africa.

The application of no-take areas in fisheries remains controversial. Critics argue that many targeted species are too mobile to benefit from area protection and that no-take areas are only appropriate for resident species. The degree of protection does not depend on the size of the no-take area but rather on the time fish reside inside its boundaries during key life-history events (i.e., spawning) and during periods of peak fishing activity. We evaluated the potential of a small no-take marine protected area (MPA) inside a coastal embayment as a harvest refuge for a mobile, possibly migratory, long-lived fish species. We used acoustic telemetry to track movements of 30 transmitter-tagged white stumpnose (Rhabdosargus globiceps) across and on both sides of the boundary of a small (34 km(2)) no-take area over a full year. Being landlocked on 3 sides, the location of the MPA inside the lagoon made it practical to detect all boundary crossings and to calculate the time individual fish used the MPA. We detected frequent movements across the boundary, with strong seasonal and individual variations. There were significant differences in MPA use patterns between fish from different release areas. The time spent in the MPA by individual fish during summer (mean 50%; max 98%) was out of proportion with the size of that area (4% of total habitat). Summer coincided with peak recreational fishing activity and with the spawning season of this species. The small MPA provided a refuge for a part of the spawning stock of white stumpnose. Our findings suggest that if strategically placed, a small no-take area can be effective in protecting mobile species and that models of spillover from no-take areas should account for seasonal and individual variation in area use and the spatiotemporal distribution of fish and fishers.